Mitochondrial phylogenomics supports a Carboniferous origin of Xenonomia

Polyneoptera includes some of the best-known insect species, such as grasshoppers and cockroaches. While the evolutionary history of many Polyneoptera orders has been thoroughly explored, others have been partially overlooked. This is the case with Xenonomia, a clade consisting of two species-poor insect orders with a relatively recent taxonomic history: Mantophasmatodea and Grylloblattodea. Here, we provide a temporal framework for their evolution, leveraging a mitochondrial phylogenomics approach encompassing all Polyneoptera orders. To strengthen the confidence in our divergence times estimation, we specifically focused on the possible impact of phylogenetic biases, such as long branch attraction, the influence of specific fossil priors, the use of nucleotide or amino acid alignments, and different clock models. Our results consistently support the origin of Xenonomia during the Carboniferous, and the divergence between the two orders is inferred to have happened before the Permian. While Grylloblattodea diversification is inferred to have occurred earlier than that of Mantophasmatodea, extant species of both orders most likely diversified after the Permian/Triassic and Triassic/Jurassic mass extinctions. Our molecular divergence time analyses complement the fossil record and support the ancient relict status of these two polyneopteran orders.